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A Century of Biodiversity: Some Open Questions and Some Answers Biodiversity ISSN: 1488-8386 (Print) 2160-0651 (Online) Journal homepage: http://www.tandfonline.com/loi/tbid20 A century of biodiversity: some open questions and some answers Roberto Cazzolla Gatti To cite this article: Roberto Cazzolla Gatti (2017) A century of biodiversity: some open questions and some answers, Biodiversity, 18:4, 175-185, DOI: 10.1080/14888386.2017.1407257 To link to this article: https://doi.org/10.1080/14888386.2017.1407257 Published online: 08 Dec 2017. Submit your article to this journal Article views: 91 View related articles View Crossmark data Full Terms & Conditions of access and use can be found at http://www.tandfonline.com/action/journalInformation?journalCode=tbid20 BIODIVERSITY, 2017 VOL. 18, NO. 4, 175–185 https://doi.org/10.1080/14888386.2017.1407257 A century of biodiversity: some open questions and some answers Roberto Cazzolla Gatti Biological Institute, Tomsk State University, Russia ARTICLE HISTORY Received 2 March 2017; Accepted 16 November 2017 Introduction Some (partially or completely) resolved questions about biodiversity The study the biodiversity is not just an attempt to under- stand the differences ro similarities between species, habi- (a) What is biodiversity? tats or genomes. It also includes an understanding of how Most ecological models that try to explain biodiversity are nature regulates the processes that characterise ecosystems too simplistic and are often considered outdated (Cazzolla and ensure their functionality. It means as well to increase Gatti 2011; Connell 1980). Moreover, recently, the prin- the ability to predict the impact of current and future ciples based on competitive exclusion (Gause 1934) for anthropogenic actions on nature (Midgley et al. 2002). the explanation of biodiversity have been criticised from Biodiversity is what makes the Earth a unique planet. both theoretical and empirical approaches (Cazzolla Gatti In the Universe, for all we know, or at least in our galaxy, 2016a). life is the exception rather than the rule (Dick 1999). It has been, therefore, suggested that niches are impor- And even if a day when we are able to document the tant to attain a genetic equilibrium that favours species presence of living beings on other planets will come, (Levine and HilleRisLambers 2009) and facilitates the cre- this will also be an exception. The appearance of life is a ation of new niches (Gatti, Hordijk, and Kauffman 2017a). phenomenon that has always fascinated mankind since What has emerged is that to reach a thorough understand- without it, our human species would have never existed ing of the evolutionary dynamics of biodiversity, which and no hairless biped would have studied life, with great could somehow explain the current distribution patterns admiration, and destroyed it with equal force (Cardinale and mechanisms of coexistence, we must consider both et al. 2012). the biogeographic and phylogenetic approaches (Cazzolla Species themselves are a mystery like the very existence Gatti 2016b; Pimm et al. 2014). of life (Wiley 1978). Why has evolution led to the ‘endless beautiful forms’, as Charles Robert Darwin called them, (b) Why are there so many species in the world? instead of letting one, amazingly adapted, dominate the evolutionary line? The idea that life needs other life forms in order to con- In recent years the advancement of ecological sciences tinue to exist, and thrive, has been frequently stressed in allowed a better understanding of some fundamental bio- the last years (Bastolla et al. 2009; Cazzolla Gatti 2011). diversity patterns (Gaston 2000). In less than a few dec- Thanks to species by-products which emerged when the ades most of the answers to the key issues about biological first living organism appeared on Earth this has certainly diversity have been enriched with new hypotheses and favoured the evolution of other species (Wilkinson 2006). theories (Ricklefs 2004). However, many other questions Over time, biodiversity emerged (Cazzolla Gatti 2016a; are still open. 2017a; Guill and Drossel 2008). CONTACT Roberto Cazzolla Gatti [email protected] This article was originally published with errors. This version has been corrected. Please see Corrigendum (https://doi.org/10.1080/14888386.2017.1416756). © 2017 Biodiversity Conservancy International R. CAZZOLLA GATTI 176 A new formulation of the ecological niche concept – that the species themselves allow the enhancement of which no longer sees a niche as a static set of conditions niches in terms of space and number. Each species plays and rules referred to any species, but rather as a dynamic a fundamental role in facilitating the colonisation of the evolution of potential and unprestatable ‘biological spaces other species, by simply modifying the basic environ- dependent on the species diversity’ – has been proposed ment and exponentially increasing the available niches’ (Cazzolla Gatti 2011). This network of relationships is dimension (Cazzolla Gatti 2011). Niches emergence is enriched with new connections each time a node is pro- autocatalytically facilitated by the presence of biodiver- duced by the network (i.e. through facilitation, autopoiesis sity itself. The fact that biodiversity can be considered and autocatalysis; Gatti, Hordijk, and Kauffman 2017a). as a system of autocatalytic sets (Gatti, Hordijk, and Kauffman 2017a) offers a possible answer to the fun- damental question of why so many species can coexist (c) What are the main drivers of biodiversity in the same ecosystem. evolution? After a critical review of the different kinds of speci- (e) Can we explain biodiversity global gradients? ation, over the years many authors have attempted to understand why the competitive exclusion principle For over two centuries, scientists have tried to disentan- has long been considered as a general low to explain gle the spatial patterns of fine-scale and global biodiver- biodiversity (Connell 1980; Tilman 1994; Vandermeer sity gradients in order to understand how biodiversity is et al. 2002). Some of them argued that the likely reason distributed on the planet (Rahbek 1995; Wallace 1878). lies in the fact that ecologists have not questioned some Given that some places on Earth are home to more spe- of the principles of natural selection. A new conceptual cies than others, an important biogeographical property of model was recently proposed (Cazzolla Gatti 2016a). It biodiversity is the latitudinal gradient of species richness. was developed in an attempt to unify the new trends in Many hypotheses have been proposed – often empirically biodiversity and evolutionary science. It argues in favour tested – to become real theories (Hillebrand 2004; Kraft of the importance of avoidance of competition, coop- et al. 2011; Wilson and MacArthur 1967) in order to eration, biological history, symbiosis (endosymbiosis explain this distribution gradient. These have been divided and endogenosymbiosis) and three-dimensionality, as into two broad categories: biogeographical/evolutionary the main forces that structure the biological diversity and ecological hypotheses. Almost all of them are, how- of ecosystems. ever, not mutually exclusive and, at the same time, do not completely explain this gradient (Currie et al. 2004; Jansson and Davies 2008). (d) How is biodiversity arranged in space? However, one of the most corroborate ecological and Ecologists argued that the exceptional diversity of species evolutionary hypotheses (Hutchinson 1959; Pianka 1966) on Earth is, at least, partly due to the different ways in is that climate (C) influences net primary production which species that are potential competitors have evolved (NPP) in such a way that, by raising/lowering the availa- traits that make them specialised and able to partition the ble biomass (B), and so increasing/decreasing the number exploitation of resources (Chesson 2000; Finke and Snyder of individuals (I), it ‘controls’ the number of species (S). 2008; Loreau and Hector 2001; Schoener 1974). However, So far, the C→NPP→I→S causal relationship has not been it has been also suggested that biodiversity is autopoi- clearly demonstrated because it does not seem mediated etic (Cazzolla Gatti 2016a; Fath et al. 2007; Ulanowicz directly by biomass (Currie et al. 2004). In other words, 1998) and autocatalytic (Gatti, Hordijk, and Kauffman there is no reason to believe that areas with higher biomass 2017a), because it is a system able to produce more of its would also have higher biodiversity. It could also be that own complexity than that produced by its environment. a few species with many individuals accounts for most of Biodiversity (as an autopoietic and autocatalytic system) the biomass in the ecosystems. requests more energy (solar light) and dissipates more By suggesting that the latitudinal gradient of biodiver- entropy (waste disordered energy) than a not-autopoietic/ sity can be, instead, explained as differences in the fractal not-autocatalytic system. dimension of the ecosystem niches, it has been shown The biodiversity-related niches differentiation theory that it is not the number of individuals or the biomass (BNDT) has been proposed to stress that the process of that influence species richness, but rather the extent of niches differentiation is strictly addressed and driven climate and primary production on niche variables. In by species (Cazzolla Gatti 2011). This theory suggests other words, it has been proposed that biodiversity, in that the number of niches in an ecosystem depends on addition to
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